Shell Shape Variation in Populations of Common Cockle Anadara Oceanica
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Biodiversity Journal, 2020,11 (3): 703–715 https://doi.org/10.31396/Biodiv.Jour.2020.11.3.703.715 Shell shape variation in populations of common cockle Anadara oceanica (Lesson, 1831) (Bivalvia Arcidae) from the intertidal areas of Margosatubig, Zamboanga del Sur (Philippines) Ranjiv D. Alibon1*, Alea Ester T. Ordoyo1, Jessa Mae P. Gonzales1, Melbert C. Sepe1, Mark Anthony J. Torres3 & Genelyn G. Madjos1,2 1Department of Biological Sciences, College of Science and Mathematics, Western Mindanao State University, Zamboanga City, Philippines 2Research Utilization, Publication and Information Dissemination Office, Western Mindanao State University, Zamboanga City, Philippines 3Department of Biological Sciences, College of Science and Mathematics, Mindanao State University - Iligan Institute of Technology, Iligan City, Philippines *Corresponding author, email:[email protected] ABSTRACT The advent of geometric morphometrics opened an area to study morphological variations in organisms. Thus, the aim of this study is to use outline-based geometric morphometrics to de- scribe variations in the shell shapes of the left and right valves of Anadara oceanica (Lesson, 1831) (Bivalvia Arcidae) populations from the two neighbouring intertidal zones of Margosat- ubig, Zamboanga del Sur, Philippines. Herein, there were two levels of analyses that were em- ployed: first, the shell shapes of the outer left and right valves between populations were compared; second, the shell shapes within population were quantitatively determined in terms of its symmetry. Results revealed significant variations both in the left and right valves of A. oceanica between populations. The variations observed are characterized by the deformations in the umbonal and anteroventral angles and in the dorsal, anterior and ventral margins of the outer shell both in the left and right valves. Although further studies are necessary in order to elucidate these variations, the second analysis revealed that the detected asymmetry in the shell shapes within A. oceanica populations was the cause of variation within populations that con- tributed to the significant variations between populations. Considering that the two sites are not geographically isolated, the results herein clearly proved that shell shape variation could also occur in neighbouring populations. The variations in the shell shapes of A. oceanica pop- ulations may have implications to habitat adaptation which aid in understanding the nature of this species especially those dwelling in the intertidal areas of Margosatubig, Zamboanga del Sur, Philippines. KEY WORDS Asymmetry; environmental conditions; geometric morphometrics, habitat adaptation; neighbouring populations. Received 01.04.2020; accepted 26.05.2020; published online 18.09.2020 INTRODUCTION shells with a heavy periostracum and taxodont hinge. It is commonly called as cockle as it morpho- The common cockle, Anadara oceanica (Les- logically resembles the European cockle Cerasto- son, 1831), = maculosa (Reeve, 1844), (Bivalvia Ar- derma edule (Linnaeus, 1758) (Bivalvia Cardiidae) cidae) is characterized by its trapezoidal ribbed which in fact belongs to the family Cardiidae. Thus, 704 RANJIV D. ALIBON ET ALII this terminology does not have any taxonomic sig- the traditional method because it effectively avoids nificance. Further, it is also called blood clam be- confusion between size and shape by preserving the cause of its specialised invariable occurrence of red shape variables and the main geometric properties blood pigments haemoglobin (Davenport & Wong, of the samples (Webster & Sheets, 2010). 1986). It typically occurs in habitats ranging from It is hypothesized that shell shape variation is an the intertidal zone on wave exposed sandy shores to adaptive strategy of bivalves in response to its cur- the marginally subtidal areas of sheltered mudflats rent ecological conditions (Alibon et al., 2018). up to the higher tidal levels within the mangrove Thereby, shell shape variations in A. oceanica pop- areas and even extend into deeper subtidal areas ulations may have implications to habitat adaptation (Brotohadikusumo, 1994). Due to the absence of which can help in understanding the nature of this well-developed siphons, A. oceanica is considered species especially those dwelling in the intertidal as a poor burrower, meaning it cannot delve in the zones of Margosatubig, Zamboanga del Sur, Philip- substrate at any depths (Brotohadikusumo, 1994). pines. Accordingly, pollution of marine water caused This poor burrowing behaviour of A. oceanica by improper disposal of residential wastes and rural makes it highly exposed in receiving high concen- run-offs is one of the main ecological concerns in trations of natural and anthropogenic wastes in the this locality and that the presence of A. oceanica in intertidal zones such as inorganic and organic nutri- this area suggests a tolerance to the current ecolog- ents, soil and sediments, and pollutants (Sithik et al., ical conditions that could be influencing its shell 2009). Just like other bivalves, A. oceanica has lim- shape. Thus, this study was conceptualized with the ited mobility that restricts its ability to avoid adverse aim to describe variations in the shell shapes of A. conditions due to its sedentary behaviour (Sharma oceanica into two levels of analyses. First, the shell et al., 2016). Hence, populations of this species are shapes of the outer left and right valves between the good candidates for the detection of different types two different A. oceanica populations from Mar- and levels of stress. gosatubig were compared using outline-based geo- Due to the fact that shell is the most variable part metric morphometrics. Second, the shell shape of A. of a bivalve species and is largely affected by envi- oceanica within population was scored for differ- ronmental conditions (Uba et al., 2019), it has been ences in shapes between its left and right valves, oth- the most widely used part in studying Anadara erwise known as fluctuating asymmetry which is a species, specifically by focusing on shell shape potential bioindicator of environmental stress in variation. Succeeding studies in Anadara species populations (Trono et al., 2015). supported significant shell shape variation within and between populations (Mzighani, 2005; Faulkner, 2010; Lodola et al., 2011; Finogenova et MATERIAL AND METHODS al., 2013; Souji & Radhakrishnan, 2015; Aydin et al., 2014; Qonita et al., 2015; Meshram & Mohite, Description of sampling sites and collection of 2016). These studies, among others, focused on tra- samples ditional approaches in morphometric studies (i.e., analysis of linear distances). However, these meth- A total of 60 adult A. oceanica individuals with ods have some statistical disadvantages such as the a similar size range of 40–50 millimeter shell length difficulty in acquiring size-free shape variables were handpicked purposively from each of the two from individuals as these measurements are highly neighbouring intertidal areas in the municipality of correlated with size (Morais et al., 2014). With this, Margosatubig, province of Zamboanga del Sur, in it is pertinent to search an applicable tool that can the Philippines; Tulog-bato, Barangay Tiguian reliably analyze shape variability and the advent of (7°34’N, 123°10’E) and Samboang, Barangay geometric morphometrics (i.e., outline-based anal- Poblacion (7°35’N, 123°10’E) shown in figure 1. ysis) has come as its solution. This quantitative tool Margosatubig is bounded on the north by the Mu- is used to determine and compare morphological nicipality of Lapuyan, on the east by the Munici- shape variations of biological structures (Sansom, pality of Dimataling, on the west by the 2009). Thus, this method was employed in this Municipality of Malangas and on the south by the study and is particularly advantageous compared to Municipality of Vincenzo Sagun. Shell shape variation in common cockle Anadara oceanica from Margosatubig, Zamboanga del Sur (Philippines) 705 Figure 1. Map of Margosatubig, Zamboanga del Sur (Philippines) showing the location of the two sampling sites. Margosatubig is one of the coastal municipali- bato whereas in higher energy regimes with ties of Dumanquillas Bay and Igat Bay in the stronger currents and moderate wave action, the province of Zamboanga del Sur which in a way it flats are generally composed of courser embodies dynamic ecosystems such as a mangrove sandy/muddy sediments similar to that in Sam- estuarine ecosystem (Tulog-bato, Barangay Tigu- boang. The taxonomy follows WoRMS (2020). ian) and a residential intertidal area (Samboang, Barangay Poblacion). The coastline in Tulog-bato Preparation and imaging of samples is characterized by mudflats and muddy shores with thick patches of mangroves that are partially en- The samples were cleaned off from their soft tis- closed with coastal body of water formed where sues before the shells were sun-dried. The umbonal freshwater from the upper land meets with saltwa- angle is distinguished clearly in the outer valve ter, a characteristic of a mangrove estuarine ecosys- (Fig. 2). The left and right valves are identified in tem. Contrarily, the coastline in Samboang is the inner valve based on the position of its pallial characterized by tidal flats and sandy shores and is sinus wherein the left valve’s pallial sinus curves to dominated with residential houses where gravels, the left and the right valve’s pallial sinus curves to coarse and sandy sediments are readily observed the